Brood Success of the Mud-Daubing Wasp Sceliphron Jamaicense (Hymenoptera: Sphecidae) in a Desert Environment
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Revista Mexicana de Biodiversidad Rev.Mex.Biodivers. 89 (2018): 466-470 Ecology Brood success of the mud-daubing wasp Sceliphron jamaicense (Hymenoptera: Sphecidae) in a desert environment Éxito de anidación de la avispa lodera Sceliphron jamaicense (Hymenoptera: Sphecidae) en un ambiente desértico Christopher K. Starr a, *, Armando Falcón-Brindis b, María Luisa Jiménez b a Department of Life Sciences, University of the West Indies, St Augustine, Trinidad & Tobago bLaboratorio de Aracnología y Entomolología, Centro de Investigaciones Biológicas del Noroeste, S.C. Instituto Politécnico Nacional 195, Playa Palo de Santa Rita Sur, 23096 La Paz, Baja California Sur, Mexico *Corresponding author: [email protected] (C.K. Starr) Received: 03 August 2017; accepted: 21 January 2018 Abstract Examination of old Sceliphron jamaicense (Fabr.) nests at 4 localities near the tip of the Baja California peninsula showed levels of brood success (i.e., the fraction of fully provisioned, closed cells producing viable adult offspring) between 24.6% and 58.1% per site. Failure was predominantly at the pupal stage at all localities. Successful cells are commonly reutilized by other solitary wasps, primarily Trypoxylon (Trypargilum) dubium Coville and Chalybion californicum (Sassure, 1867). There is a pronounced clumping tendency in successes and failures among nests. Keywords: Brood success; Sceliphron jamaicense; Sphecidae; Araneae; Baja California Sur; Mexico Resumen Se llevó a cabo la revisión de nidos de Sceliphron jamaicense (Fabr.) en 4 localidades hacia el extremo austral de la península de Baja California. Las crías de las celdas mostraron niveles de éxito variable (i.e., la proporción de celdas cerradas y provistas con arañas que contenían adultos viables), entre 24.6% y 58.1%, según la localidad. Los fracasos se presentaron generalmente en estado de pupa. Las celdas exitosas a menudo son reutilizadas por otras avispas solitarias, sobre todo por Trypoxylon (Trypargilum) dubium Coville y Chalybion californicum (Sassure). Los éxitos y fracasos de la progenie demostraron un patrón de agregación entre los nidos. Palabras clave: Éxito de anidación; Sceliphron jamaicense; Sphecidae; Araneae; Baja California Sur; México Introduction cluster (Iwata, 1976). Under sheltered conditions, the distinctive nests presumably persist virtually unchanged Sceliphron Klug, 1801 is a worldwide genus of 35 for years (Fye, 1965). The mother wasp stocks the cells known species (Pulawski, 2016), which are relatively with paralyzed spiders as brood food (Coville, 1987). The uniform in their morphology and habits (Bohart & Menke, few species whose prey arrays have been studied each hunt 1976; Camillo, 2002). All studied species make self- a variety of spiders of different ecological types (Elgar & standing nests of mud, each of several cells in a tight Jebb, 1998; Muma & Jeffers, 1945). In Mexico, the lucae ISSN versión electrónica: 2007-8706; Universidad Nacional Autónoma de México, Instituto de Biología. Open Access bajo la licencia CC BY-NC-ND (4.0) https://doi.org/10.22201/ib.20078706e.2018.2.2416 C.K. Starr et al. / Revista Mexicana de Biodiversidad 89 (2018): 466-470 467 https://doi.org/10.22201/ib.20078706e.2018.2.2416 form of S. jamaicense occurs in Baja California and from Las Cuevas, 23°32’ N, 109°41’ W, and d) Santiago, 23°29’ mid to northern Pacific Coast (Van der Vecht & Van N, 109°43’ W. The predominant vegetation throughout the Breugel, 1968). It is known to provision with up to 36 region is tropical dry forest with 645 taxa of vascular spider species from 14 families (Jiménez et al., 1992). plants (5.7% are endemic). Mimosa lagunensis, Erythrina When a cell is stocked with spiders and her egg, flaveliformis, Acaciella goldmanii, Jatropha cinerea, the mother wasp seals it with mud and has no further Bursera microphilla, and Lysiloma microphyllum are the interaction with it except often to apply a covering layer most important species in this vegetation community. of mud up to about 3 mm thick to the nest as a whole The climate is warm steppe, semiarid BSo at 500-700 m (Bohart & Menke, 1976). Such a completed cell is not elevation and BS1 at 700-1,550 m (León-de la Luz et al., certain to yield an adult offspring. Sceliphron. jamaicense, 2012). The soils are derived from granite rocks with active like other solitary aculeate Hymenoptera, is subject to a erosion and surface rockiness. They are poorly developed number of nest parasites (Coward & Matthews, 1995; soils of sandy loam-textured slightly acid pH but poor in Crawford, 1982). These can reduce brood success —the organic matter (Maya-Delgado, 1988). fraction of completed cells from which adult offspring We collected nests on and in buildings at all localities, emerge— substantially. In addition, there is reason to although at Las Cuevas most nests were from a high suspect that the hard cell walls and covering layer can sandstone cave about 40 m in depth. We examined nests at sometimes prevent an adult offspring from emerging, the Centro de Investigaciones Biológicas del Noroeste, La further reducing brood success. O'Neill's (2001: Table 7- Paz, dismantling them systematically and recording each 3) review of brood success in 29 species of solitary wasps cell as either successful or failed with respect to brood showed a median value of 51%, with most species between production. Success was evidenced either by a species- 45% and 60%. These included 4 species of Sceliphron characteristic emergence hole (either 4 or 5 mm diameter, with success rates of 57-64%. almost always through the cell closure) or reutilization An emerging Sceliphron offspring leaves behind an by C. californicum or T. (Trypargilum) dubium, neither open, hard-walled cylinder of about 9 mm inner diameter of which is known in any failed Sceliphron cell. C. and volume 1.5 ml. While no Sceliphron species is known californicum covers its nest closure with a contrasting white to reutilize its own or conspecific cells, some other layer of uric acid broader than its own emergence hole aculeates take advantage of such ready-made brood cells, (Gess & Gess, 1980, Jayakar & Spurway, 1963) (Fig. 1). a habit known as renting (Iwata, 1964; Spurway et al., We interpreted as a failed cell one that was closed 1964; Ward, 1970, 1971). Those of S. jamaicense in Baja with a clear indication of offspring death inside. A mass California Sur are commonly reutilized by 2 other wasps: of dead, unconsumed spiders indicate that the offspring C. californicum and T. (Trypargilum) dubium (Gadar, died as an egg or larva. Death at the pupal stage was seen 2009; Jiménez et al., 1992). in a closed pupal case, often filled with the remains of a While it is relatively easy to find Sceliphron nesting great many minute parasitoids (presumably Melittobia sp.; sites, the search for active nests is much more laborious Camillo, 2002; Jiménez et al., 1992) or their pupal exuviae for humans and presumably for nest parasites. A parasite and a very small emergence hole through the cell wall. that finds 1 cell at a suitable stage for exploitation will A very few offspring reached adulthood and then were presumably find others at the same time. This leads to the unable to emerge from the brood cell. We disregarded the prediction of a clumped distribution of brood successes few cells made of concrete, evidently from construction and brood failures among nests (Hunt, 1993). sites, as well closed cells that were entirely empty, on the Our purpose here is to assess brood success of S. assumption that these latter had never been provisioned. jamaicense in a desert environment of the Cape Region of From successful cells we recorded reutilizations by C. Baja California Sur, Mexico, with emphasis on variation californicum and T. (Trypargilum) dubium. Reutilization among localities, and among nests within localities. We by C. californicum is evidenced by layer of white material predicted a highly clumped distribution of successes and on and sometimes around the cell top. The much smaller failures, such that if a parasite breached 1 cell it would T. (Trypargilum) dubium divides the cell into 2 or (more likewise breach most of the others in the same nest. commonly) 3 chambers, each serving as a brood cell, by means of transverse mud partitions, remnants of which Materials and methods persist. Occasionally a cell successfully reutilized by the former is then reutilized by the latter, although apparently During June 2016, we collected old S. jamaicense nests never the reverse. In such a case, we recorded the cell at 4 localities in Baja California Sur: a) El Triunfo, 23°48’ as reutilized by C. californicum only. It is notable that N, 110°06’ W; b) San Bartolo, 23°45’ N, 109°51’ W; c) at Las Cuevas the C. californicum-reutilized nests were 468 C.K. Starr et al. / Revista Mexicana de Biodiversidad 89 (2018): 466-470 https://doi.org/10.22201/ib.20078706e.2018.2.2416 nests in partial darkness. In the cave at Las Cuevas, we found abundant nests from near the cave mouth to all parts light enough for us to find our way with ease. This was the only locality where we found evidence of parasitism by cuckoo wasps (Chrysididae: prob. Chrysis sp.). We examined a total of 2,234 cells from at least 316 old nests. Brood success varied widely among localities from 25% (El Triunfo) to 58% (Santiago) (Table 1). At all localities, brood failure was most commonly in the pupal stage (Table 2). Trypoxylon (Trypargilum) dubium was the predominant reutilizer of S. jamaicense cells, with C. californicum also reutilizing a significant minority of cells, so that few successful S. jamaicense cells remained permanently vacant (Table 2). As seen in table 3, successes and failures showed a consistent pattern of moderate clumping among nests. In each of the 4 nest sizes there were significantly more all- success and all-failure nests and fewer mixed nests than expected (X2 test, p = 0.02 for 2-cell nests, p < 0.01 for 3-, 4- and 5-cell nests).